The role of cereal growth stages and crowding in the induction of alatae in Sitobion avenae and its consequences for population growth

Abstract. 1. Alatiform nymphs of Sitobion avenae were first recorded on wheat in 1977 and 1978 when there were 0.6 aphids per tiller and reached a maximum proportion of 0.9 of the population despite a seventeen-fold difference in aphid density between years.
2. At the same aphid density per tiller there was a higher proportion of alatiform nymphs on the tillers at an advanced than at an earlier stage of plant growth.
3. Both pre- and post-natal crowding, and late stages of plant growth, were important in inducing the development of alatae in the laboratory.
4. Changes in the quality of the host plant through its previously reported effect on reproductive and survival rates, and its effect on alate induction and emigration was important in shaping the population dynamics of S.avenae on cereals.     

Mechanisms of species‐sorting: effect of habitat occupancy on aphids' host plant selection

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Xylem ingestion by winged aphids

When aphids and their host plant are incorporated in a DC electrical circuit, phloem and xylem ingestion register as separate waveforms of the electrical penetration graph (EPG) signal. Aphids are primarily phloem feeders; xylem ingestion is seldom reported but can be induced experimentally by fasting the insects in desiccating conditions. In experiments with the black bean aphid, Aphis fabae Scop., young winged (alate) and unwinged (apterous) virginoparous adults were collected from their natal host plants (broad bean, Vicia faba L.) and allowed 3-h continuous EPG-recorded access to V. faba seedlings. Several aphids (47% of both morphs) showed ingestion from phloem sieve elements. Alate aphids also showed frequent xylem ingestion (60% of individuals), but no apterous aphids exhibited this activity. The EPG technique involves attachment of a fine gold wire electrode to each insect, a process that may affect normal behaviour at the plant surface. However, when the technique was modified to monitor the stylet activities of freely-settled aphids, high levels of xylem ingestion by alates were also recorded. The results suggest that the developmental physiology of winged aphids somehow predisposes them to xylem ingestion, possibly as a result of dehydration during the teneral period. Alate aphids may reduce their weight by fasting before take-off, giving aerodynamic benefits, but making rehydration, via xylem uptake, a priority following plant contact.     

蚜虫的表型可塑性及其遗传基础

表型可塑性(phenotypic plasticity)是有机体在适应生物或非生物环境时呈现不同表型的能力,并且有遗传基础。蚜虫是农林业的重要经济害虫,易受外部环境因素和自身遗传因素的影响而表现出表型的可塑性。本文综述了外部环境因素（如寄主植物、温度、光照、天敌等）的变异对蚜虫表型的影响。总体来说,蚜虫表型会因寄主植物的种类、品系以及发育阶段和营养状况的不同而有所差异; 温度变化对不同蚜虫种类的生殖力、生存力以及有翅蚜产生与否有极大影响。研究人员利用RAPD-PCR、微卫星等分子遗传标记确认寄主植物和温度是造成蚜虫种群遗传分化的重要因素。就内部因素而言,不同的蚜虫种类以及同一种蚜虫的不同克隆系在表型和遗传进化上也有不同程度的差异,在蚜虫受外界条件影响的不同虫态以及不同体色克隆系、不同生活周期的类群之间, 其生物学、生态学和遗传学都有所差异。分析上述各个因素对蚜虫表型可塑性的影响,对于蚜虫的生态进化研究和有效治理蚜害均有重要意义。本文在最后讨论了还有必要深入研究的诸多问题,如表观遗传调控,包括DNA甲基化、基因所在的核小体上的组蛋白的共价修饰和染色质重塑、siRNA介导的基因沉默以及微RNA（microRNA 或 miRNA）调控的基因表达变化等,又如有翅蚜的表型和遗传学研究,以及全球气候变化对蚜虫的生态进化的影响等问题。     

Evaluation of induced responses, insect population growth, and host-plant fitness may change the outcome of tests of the preference-performance hypothesis: a case study

The preference‐performance hypothesis predicts that insect preference should correspond to host suitability for offspring development. We studied the pattern of within‐plant preference in the aphid Sipha flava and its consequences for offspring performance on the host‐plant Sorghum halepense, regarding the role of induced responses of plants to aphid feeding. The consequences of within‐plant preference on aphid population growth and host‐plant traits were also evaluated. Our results showed that winged and wingless aphids preferred to settle on mature rather than young leaves. In contrast, aphid individual growth rate was higher on young leaves when compared with mature leaves, suggesting that the outcome of this test rejected the preference‐performance hypothesis. However, the inclusion of the factor ‘previous aphid infestation’ changed the outcome from a maladaptive choice to a neutral one. Thus, individual growth rates of S. flava increased when aphids developed on leaves that had been previously infested. Interestingly, aphid growth rate on previously infested leaves did not differ between young and mature leaves. On the other hand, aphid population reproductive rate was higher and the percentage of winged aphids lower when infestation occurred on mature rather than young leaves. Aphid infestation reduced plant and shoot biomass, and increased leaf mortality. These negative effects on plant traits related to plant fitness were greater when aphid infestation occurred on young leaves. Likewise, whereas infestation on mature leaves did not cause a significant reduction in the number of flowering plants compared with control plants, aphid infestation on young leaves did reduce the number of plants at the flowering stage. Consequently, if both the reproductive rate of aphids in the mid‐term, and host‐plant fitness are taken into account, the results indicate that aphid preference for mature leaves may be an adaptive choice, thus supporting the preference‐performance hypothesis.     

The role of nutrition, crowding and interspecific interactions in the development of winged aphids

1. Winged morph production in aphids is a phenotypic trait that has traditionally been seen as a response to unfavourable environmental conditions. The evidence to support this theory is reviewed and the ecological and evolutionary significance of the findings is discussed. 2. The common assertion of poor host‐plant nutritional quality leading to increased production of winged morphs does not always apply, particularly when the host‐plant quality is exceptionally poor. The available data are skewed heavily towards Myzus persicae, and for this species the dynamical change in plant quality appears to be important with respect to wing induction. 3. Crowding may be a less influential stimulus for wing induction as study methods approach natural conditions experienced by aphids on their host plant. 4. The growing evidence that interactions with other organisms can induce the production of winged morphs by aphid colonies is reviewed. In the case of natural enemies, such a response by an aphid colony may be regarded as induced defence. Wing induction may also act as a means of transmission for a virus or fungal pathogen.     

The role of refuge areas in the phenology of Rhopalosiphum padi in low rainfall cropping areas of South Australia

Extensive surveys of possible aphid habitats in South Australia indicated that irrigated perennial grass pastures in the Mount Lofty Ranges and Lower Murray Valley were summer refuges for Rhopalosiphum padi (L.) (Hemiptera: Aphididae). Large numbers of aphids build up in these pastures each year during autumn (April and May) with numbers peaking in May. The size of the May peak was related to the number of aphids surviving the summer. The proportions of alates were highest in May and August/September. Both peaks coincided with a photoperiod of between 11.2 and 11.5 h, and partial correlations suggested that aphid density, photoperiod and temperature were all significant determinants of alate production.     

Migratory tendency in aging populations of the pea aphid,Acyrthosiphon pisum

Summary Sweep samples of the aphid, Acyrthosiphon pisum, were collected from six natural populations ranging in age from one to five years. Clones were established in the laboratory from the field-collected adults and tested for their migratory tendency in two subsequent generations by measuring the percentage of winged offspring produced in response to a standard stimulus. The number of aphids in sweep samples and the percentage of winged and wingless aphids were also determined. Tests on the first laboratory generation revealed a decline in migratory tendency with the age of the source population, but no such relationship was detected in tests on the second generation. These results are consistent with an explanation based on maternal age effects and differences in adult age structure among populations of different age. They are not consistent with one based on genetic differences among populations. Older populations also had higher densities and a lower percentage of alate adults. The percentage of larvae with wing buds was positively correlated with population density, but not population age.     

棉花型和瓜型棉蚜产生有性世代能力的分化

为了明确寄主专化性是否会影响棉蚜的繁殖策略或生活史对策,采用低温和短光照组合（18℃,L∶D=8∶16）分别对棉花型和瓜型棉蚜进行有性世代的诱导,比较两寄主专化型棉蚜产生有翅蚜、雄性母、雌性母及无翅孤雌蚜的能力。结果表明：两寄主专化型棉蚜在产生有性世代能力上存在显著差异,表现为棉花型棉蚜在诱导后代中产生有翅蚜、雄性母和雌性母的比率显著高于瓜型棉蚜,并且雌、雄性母产生的时间明显早于瓜型棉蚜。瓜型棉蚜在诱导条件下产生无翅孤雌蚜的比率显著高于棉花型棉蚜,并且发现有不产生有性世代的营专性孤雌生殖个体,而在棉花型棉蚜中没有发现。产生性母蚜的棉花型和瓜型棉蚜均可同时产生孤雌胎生后代。寄主型与诱导时间长短对棉蚜有性世代的产生存在交互作用。采集于田间棉花和黄瓜上的棉蚜,也表现为棉花上的易产生有性世代,在诱导的第2代中产生性母蚜的比率显著高于黄瓜上的棉蚜,并且性母蚜比率在诱导的第2代与第3代间无显著差异; 但黄瓜上的棉蚜性母蚜产生比率第3代显著高于第2代。由此推测,棉蚜寄主专化性的形成与生活史特性的分化有关。     

Geographic and clonal variation in the milkweed-oleander aphid,Aphis nerii (Homoptera: Aphididae), for winged morph production,life history,and morphology in relation to host plant permanence

Summary Populations of the milkweed-oleander aphid,Aphis nerii, were sampled in California, Iowa and Puerto Rico. Among these localities the aphid's host plants differ greatly in permanence. I compared populations for migratory potential, measured as the proportion of winged offspring produced in response to being crowded, and for life history and morphometric traits of the subsequent adult winged aphids. I predicted a negative correlation between degree of host plant permanence and migratory potential. As predicted, aphids from Iowa, where migration on to temporary hosts must occur each year, produce a greater proportion of winged offspring (37.7%) than those from California (25.7%) or Puerto Rico (31.6%) where hosts are more permanent. However, hosts in Puerto Rico appear to be more permanent than those in California, yet the difference between populations for migratory potential was opposite to that predicted. Within California the prediction again held: aphids collected from the most impermanent sites produce the greatest proportion of winged offspring. There were no population differences for any life history or morphometric traits of winged aphids that are important contributors to fitness or migratory ability such as time to reproductive maturity, fecundity or wing length. Nor did any traits covary with migratory potential. Thus, there does not appear to be an association of life history and morphology with migratory potential that could enhance the colonizing ability of migrant aphids. I was unable to detect population differentiation for life history and morphology even though there is ample genetic variation within populations on which selection could act and an absence of constraints arising from genetic correlations that could prevent appropriate evolution of traits within populations. The exploitation of temporary host plants therefore occurs by an increase in the number of colonists produced and not by change in life history or morphology of those colonists.     

宁夏地区麦长管蚜远距离迁飞的研究

通过1992—1978年的调查试验, 初步明确了以下事实:本地越冬的麦长管蚜尚未羽化, 田间已出现有翘成蚜;小麦苗期发现了穗期才能产生的穗型蚜;在冰冻期的高山上多次捕到有翅麦蚜;有翅成蚜大范围的“同期突发”现象;外来麦蚜占春季田间有翅蚜的98.8%以上.首次提出, 在有本地蚜源的情况下, 存在外来蚜源, 且外来蚜源可以成为春季田间麦长管蚜群的主体.本文分析讨论了迁入蚜源的可能基地和须进一步研究的问题.     

The mid‐season crash in aphid populations: why and how does it occur?

Abstract. 1. Aphid populations on agricultural crops in temperature regions collapse over a few days from peak numbers to local extinction soon after mid‐summer (e.g. mid‐July in the U.K.). The populations recover 6–8 weeks later. There is anecdotal or incidental evidence of an equivalent mid‐season population crash of aphids on grasses and forbs in natural vegetation. 2. The ecological factors causing the mid‐season population crash of aphids include a decline in plant nutritional quality and increased natural enemy pressure as the season progresses. Extreme weather events, e.g. severe rainstorms, can precipitate the crash but weather conditions are not a consistent contributory factor. 3. The population processes underlying the crash comprise enhanced emigration, especially by alate (winged) aphids, depressed performance resulting in reduced birth rates, and elevated mortality caused by natural enemies. 4. Mathematical models, previously applied to aphid populations on agricultural crops, have great potential for studies of aphid dynamics in natural vegetation. In particular, they can help identify the contribution of various ecological factors to the timing of the population crash and offer explanations for how slow changes in population processes can result in a rapid collapse of aphid populations.     

Within-plant distribution of cotton aphids, Aphis gossypii Glover (Hemiptera: Aphididae), in Bt and non-Bt cotton fields

Knowledge of the vertical and horizontal distribution of Aphis gossypii Glover (Hemiptera: Aphididae) on genetically modified cotton plants over time could help optimize decision-making in integrated cotton aphid management programs. Therefore, the aim of the present study was to determine the vertical and horizontal distribution of A. gossypii in non-transgenic Bt cotton and transgenic Bt-cotton over time during two cotton seasons by examining plants throughout the seasons. There was no significant interaction between years and cotton cultivar treatments for apterous or alate aphids. Considering year-to-year data, analyses on season-long averages of apterous or alate aphids showed that aphid densities per plant did not differ among years. The number of apterous aphids found per plant for the Bt transgenic cultivar (2427 apterous aphids per plant) was lower than for its isoline (3335 apterous aphids per plant). The number of alate aphids found per plant on the Bt transgenic cultivar (12.28 alate aphids per plant) was lower than for the isoline (140.56 alate aphids per plant). With regard to the vertical distribution of apterous aphids or alate aphids, there were interactions between cotton cultivar, plant age and plant region. We conclude that in comparison to non-Bt cotton (DP 4049), Bt cotton (DP 404 BG (Bollgard)) has significant effects on the vertical, horizontal, spatial and temporal distribution patterns of A. gossypii, showing changes in its distribution behaviour inside the plant as the cotton crop develops. The results of our study are relevant for understanding the vertical and horizontal distribution of A. gossypii on Bt cotton cultivar (DP 404 BG (Bollgard)) and on its isoline (DP 4049), and could be useful in decision-making, implementing controls and determining the timing of population peaks of this insect.     

植物病媒昆虫的翅型分化

翅多型现象是昆虫非遗传多型性的一种表现,包括不具飞行能力的短翅型或无翅型,以及可以进行长距离迁飞的长翅型或有翅型。翅多型现象常发生在可以携带病原并将其传播给植物宿主的媒介昆虫中,对植物病害的时空分布与暴发有重要影响。本文从翅型分化的遗传规律、诱导因素、分子机制和伴随翅型分化的其他生理表现4个方面,对植物病原主要传播媒介蚜虫和飞虱的翅型分化研究进行综述和梳理。昆虫翅型分化的诱导因素主要包括温度、湿度和光周期等非生物因素以及虫口密度、宿主营养、病毒等生物因素;而其内在的分子机制大多是通过胰岛素/胰岛素样生长因子信号(IIS)通路、c-Jun氨基末端激酶(c-Jun NH₂-terminal kinase, JNK)信号通路、Wingless和嗅觉受体SaveOrco等调控。翅型分化的同时伴随着生理状态的变化,表现为短翅型具有更强的繁殖能力和长翅型含有更丰富的飞行肌结构成分。目前,昆虫翅型分化的研究尚不够完善,有许多需要解答的问题,如找到胰岛素/胰岛素样生长因子信号通路中真正发挥功能的靶基因,JNK如何调控翅型分化以及虫媒病毒影响媒介昆虫翅型的分子机理。本综述可为控制虫...     

Endophytic fungi decrease available resources for the aphid Rhopalosiphum padi and impair their ability to induce defences against predators

Abstract.  1. The production of winged morphs is a well known mechanism of induced defence in aphids to escape from natural enemies, and is also a reaction to poor resource quality.
2. Host plants of aphids often associate with endophytic fungi that have been shown to reduce the fitness of some species of aphids.
3. It was hypothesised that endophyte infection of host plants that represent a low quality plant resource should increase the aphid's induced response to a predator because both low plant quality and predator presence represent a stronger cue for wing production than predator presence alone.
4. In a laboratory experiment, bird cherry-oat aphids Rhopalosiphum padi L. were exposed to the factors predator threat and endophyte infection and the effects of these factors on the proportion of winged morphs produced by the aphid colonies was analysed.
5. The presence of endophytic fungi strongly decreased aphid colony sizes. When a predator threat was present, all colonies on endophyte-free grasses produced winged morphs whereas only a few colonies were able to produce winged morphs on endophyte-infected grasses. However, these few colonies produced larger proportions of winged morphs than colonies on endophyte-free grasses. Without a predator threat, no colonies on endophyte-infected grasses produced any winged morphs.
6. These results show that aphids in stressed conditions and with reduced fitness will only invest in inducible defences when predators are present but are unable to produce winged morphs in response to endophyte presence.     

“8455”小麦植株内含物等因子与麦蚜(长管蚜,二叉蚜)种群有翅率的关系

通过对不同肥力条件下“8455”小麦植株内含物及温湿度、平均拥挤度等因子与长管蚜、二叉蚜种群有翅率关系的研究与分析,得出,小麦植株内含物中含水量、可溶性糖含量、全氮量及相对湿度、平均拥挤度是影响长管蚜种群有翅率的主要因子;含水量、全钾量和相对湿度是影响二叉蚜种群有翅率的主要因子。     

Impact of wheat-mung bean intercropping on English grain aphid (Hemiptera: Aphididae) populations and its natural enemy

The effects of intercropping wheat, Triticum aestivum L., with mung bean, Vigna radiate L., on the populations of English grain aphid, Sitobion avenae (F.) (Hemiptera: Aphididae), and its natural enemies were evaluated by field and laboratory experiments. The population densities of aphids and their natural enemies were evaluated in the intercropped field against different row ratio combinations of wheat-mung bean. Results showed that wheat-mung bean intercropping caused a drop in aphid densities, and the ratio 12 wheat: 4 mung bean brought about the largest drop (> 8%). In addition, the population densities of coccinellids (ladybirds) and parasitoids and the species diversity of all the natural enemies of aphid were higher in the intercropped field than in the field planted only with wheat. However, intercropping did not influence the community indices (evenness and index of dominance concentration) of the natural enemies. Y-tube olfactometer bioassays were carried out in the laboratory to test whether odor blends of host and nonhost plants affect the host selection of S. avenae. Bioassays indicated that both apterous and alate aphids significantly preferred host plant odor over odor blends of host and intercropped species. Hence, the olfactory-based host location of aphids in the field might be affected by intercropping. The intercropping experiment clearly showed that increased crop species diversity suppresses aphid population growth and preserves the population of natural enemies of aphids. Our results also provide support for the "resource concentration hypothesis" and the "enemies hypothesis".     

The cost of being able to fly in the milkweed-oleander aphid,Aphis nerii (Homoptera: Aphididae)

Summary In the wing dimorphic milkweed-oleander aphid,Aphis nerii, winged aphids begin reproducing about 1.5 days after wingless aphids. The longer maturation period is primarily due to slower development since even adult eclosion by winged aphids takes place after wingless aphids begin reproducing. The delay is not due to a post-eclosion, pre-reproductive flight since, beginning with the fourth instar, larval winged aphids were reared at a density of one per plant and the vast majority were not stimulated to fly under such low-density conditions. Thus, the ability to fly incurs a fitness cost in terms of delayed reproduction, irrespective of whether flight actually occurs. We did not observe a difference between morphs for lifetime fecundity, even though wingless aphids have larger abdomens than winged aphids and for both morphs there is a significant correlation between abdomen width and fecundity. Offspring produced by wingless aphids over the first four days of reproduction are larger than those produced by winged aphids, and the size difference at birth is maintained into adulthood. However, there are no differences in life history traits between these offspring, including maturation period and lifetime fecundity. Thus, reduced body size does not increase the cost of being able to fly, at least under the conditions of these experiments. The cost of being able to fly in this species should favor reduced production of winged individuals in populations that exploit more permanent host plants.     

Interannual dynamics of aerial and arboreal green spruce aphid populations

Partial defoliation of spruce by the green spruce aphid Elatobium abietinum (Walker) is a recurrent event in European and, increasingly, North American forests. The patterns of insect abundance on trees have never been satisfactorily described by a numerical model despite considerable knowledge of endogenous and exogenous factors in the population dynamics of the species. Long-term field population estimates of the aphid on foliage provided the opportunity to evaluate such a model. Unlike comparable models for tree-dwelling aphids, this was also applicable to almost completely independent aphid field data derived from the Rothamsted Insect Survey’s nationwide network of suction traps. Although based on relatively few parameters, the model was robust in its predictions of alate aphids geographically remote from the forest in which the original population was estimated. The population maximum, which causes the greatest forest damage, is reached in early summer and can be predicted from knowledge of winter temperature (chill bouts), spring temperature (thermal sum), and interannual negative feedback (density dependence). The model provides confirmation that alate populations of spruce aphids, upon which a number of other extensive studies have been based, are ultimately influenced by similar endogenous and climatic factors and that they are a reasonable proxy for aphids on trees.